CHEMICAL ECOLOGY – Chemical Signals as Attractants, Repellents and Aggregation Stimulants - Baldwyn Torto CHEMICAL SIGNALS AS ATTRACTANTS, REPELLENTS AND AGGREGATION STIMULANTS Baldwyn Torto International Center of Insect Physiology and Ecology, Kenya Keywords: Semiochemical, kairomone, allomone, synomone, pheromone, attractant, repellent, aggregation stimulant, receptors, herbivores, carnivores, predators, parasitoids, tritrophic, co-evolution Contents 1. Introduction 2. Classification of Chemical Signals 3. Detection and Decoding of Semiochemicals 4. Approaches to the Isolation and Identification of Semiochemicals 5. Attractants in Insect Communication 5.1 Plant-produced attractants 5.2 Vertebrate-produced attractants 5.3 Sex attractants 6. Repellents 7. Aggregation stimulants 8. Semiochemicals and Parsimony 9. Practical Applications 10. Conclusions Acknowledgements Glossary Bibliography Biographical Sketch Summary Chemical signals in the environment mediate behavioral and/or physiological interactions between different organisms. A chemical signal in the environment may benefit the emitter,UNESCO the receiver or both, a nd– may EOLSS evoke a behavioral response such as attraction, repellence or aggregation, responses that are dependent on the nature, quality and quantity of the chemical signal emitted. 1. IntroductionSAMPLE CHAPTERS In nature, organisms regularly encounter signals from their environment, signals that may be visual, auditory, physical or chemical, and which influence their behavior. Chemicals in the gaseous phase constitute the odor signals that are conveyed in the air, and are detected by their receivers as a coded message. A coded chemical message may be simple or complex, but when decoded, it may benefit its emitter, the recipient, or both. The message may evoke in the recipient an immediate short-term and reversible behavioral response (releaser effect), or long-term irreversible physiological and/or biochemical changes (primer effect). The latter mainly occur in the endocrine system of ©Encyclopedia of Life Support Systems (EOLSS) CHEMICAL ECOLOGY – Chemical Signals as Attractants, Repellents and Aggregation Stimulants - Baldwyn Torto the organism. These types of chemical signals are released in minute quantities, in most cases far below human detection by smell. While many of these chemical signals have been discovered mediating interactions among various animal taxa, most of the research has focused on the interaction among insects. An important reason for this is that insects constitute the largest group of animals on earth, with a large number of species being economically important as pollinators, crop pests and vectors of animal and human diseases. Insects exhibit many forms of behavioral patterns, however, this article focuses on chemical signals that evoke three types of short-term behavioral responses in insects: attraction, repellence and aggregation stimulation. 2. Classification of chemical signals A chemical signal that conveys a message between organisms of the same or different species is referred to as a ‘semiochemical’ (Gk. semeon, sign or signal). Some researchers refer to these types of chemical signals as infochemicals since they only convey information in order to distinguish them from venoms and toxins used specifically by certain organisms for defense. Semiochemicals mediate diverse natural interactions involving terrestrial and aquatic organisms. Among insects, interactions with plants, animals, and microbes are recognized. Their sources of production and emission are equally diverse: flowers, leaves, stem bark and roots, microbial secretions, insect body glands, saliva, reproductive organ fluids and excretory products. Semiochemicals are mainly secondary metabolites, produced as by-products from metabolic processes that occur in the tissues of organisms or from enzymatic or microbial activity on ingested food. Although they vary in complexity, semiochemicals that are conveyed in the gaseous phase generally comprise low molecular weight compounds (less than 250 M.W.). Four main classes of semiochemicals are recognized: Kairomone- Gk. kairos (= opportunistic), a message-bearing chemical that provides an adaptive advantage to the receiver (e.g., an attractant or aggregation stimulant). Allomone– Gk. allos (= other), a message-bearing chemical that coveys an adaptive advantage to the emitter (e.g., repellent). Synomone- Gk. syn (= with), a message-bearing chemical that conveys a mutual advantage to both the emitter and the receiver. (e.g., floral odor attracts pollinatingUNESCO insects, for example honeybee, – EOLSS the flowering plant benefiting from pollination by the insect that in turn is rewarded with nectar or pollen). Pheromone- Gk pherein (= to transfer; horman = to excite), a message-bearing chemical emitted and received by an organism of the same species. SAMPLE CHAPTERS Using biological assays, which often involve the application of the chemical to a neutral material, it is possible to discriminate between semiochemical types. For example, an attractant chemical will elicit a positive displacement (i.e., movement) of the recipient organism towards it, while a repellent chemical will elicit a negative displacement of the recipient away from it. On the other hand, an aggregation stimulant will draw individuals of the same species together irrespective of their sexes to group or aggregate. Semiochemicals are usually not restrictive in their function and may exert different effects depending upon the insect species or conditions under which they are presented. For example, a semiochemical released at a given dose may on the one hand ©Encyclopedia of Life Support Systems (EOLSS) CHEMICAL ECOLOGY – Chemical Signals as Attractants, Repellents and Aggregation Stimulants - Baldwyn Torto function as an attractant for one insect species, while on the other, it may function as a repellent or aggregation stimulant in another species. It may for the same individual insect function as an attractant or repellent at different doses, or evoke no response at all if presented as a single component when activity is due to a blend of components. In the majority of studies, semiochemicals have been found to be composed of blends rather than single components. However, for even simple blends, optimal activity is achieved when the precise ratios of individual components in the synthetic blend simulate the natural situation. 3. Detection and decoding of semiochemicals Insects detect chemical signals in the environment via sense organs that are endowed with specific receptors, called ‘chemoreceptors’. There are two types of chemoreceptors; those that detect chemicals in solution form which are denoted as taste or gustatory receptors, and those that detect chemicals in gaseous form, referred to as olfactory receptors. These receptors are housed in minute or microscopic hair-like structures (setae), which are distributed on various appendages located on different parts of the body. Chemoreceptors can detect and recognize the individual components of a chemical signal in a ‘lock and key’ mode (i.e., receptor-substrate interaction). While gustatory chemoreceptors are largely concentrated on the mouthparts (maxillary and labial palps, labrum), tarsi, and ovipositor of insects, a large population of the olfactory receptors is located on the antennae. In olfactory reception, an insect exposed to an odor source first detects the chemical signal using its antennae, which involves recognition of the active components in the signal by chemoreceptors according to their structures, or loosely, according to their ‘sizes and shapes’ or spatial arrangement (enantiomeric composition). These chemoreceptors then relay messages through the antennal nerve to the central nervous system (brain), which decodes the messages via cellular and metabolic activity. This may in turn activate sets of muscles whose reaction may manifest as a behavioral response. The response may be an orientation and/or a displacement towards or away from the source of the chemical signal to signify in the recipient of the chemical signal the presence of an attractant or a repellent, respectively. The accuracy with which a chemical signal is decoded is dependent on the nature, quality and quantity of the signal. In an evolutionary sense, most chemical messages are interpreted in an associative context i.e., as an indicator of an available resource e.g. food, a mate,UNESCO or a suitable egg-laying site. They– mayEOLSS also denote the presence of other organisms in the environment that may either be beneficial, or harmful to the recipient. 4. Approaches to isolation and identification of semiochemicals SAMPLE CHAPTERS The message content of a chemical signal is often complex, composed of a range of chemical compounds. An insect exposed to the chemical signal selectively responds to the active components in the signal. The active components in the chemical signal can be isolated and identified using interdisciplinary approaches involving insect behavior, sensory physiology, analytical and organic chemistry, biochemistry and ecology. Recent years have seen breakthroughs in analytical techniques, which now permit for rapid screening of semiochemicals in more sensitive bioassays and their isolation and identification from relatively smaller amounts of material. There are several non- destructive methods available for collecting natural odors from living organisms, which ©Encyclopedia of Life Support Systems (EOLSS) CHEMICAL ECOLOGY – Chemical